The Mango Botany Production and Uses

CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses(ed. R.E. Litz) 1

1 Introduction: Botany and Importance

S.K. Mukherjee1 and R.E. Litz21Calcutta University, Calcutta, India2University of Florida, Florida, USA

1.1 Introduction 11.2 Description of Mango 2

The tree 2Flowers 2The fruit 3The seeds and polyembryony 4

1.3 History of Cultivation 5Origin of Mangifera indica 5Domestication of mango 9Distribution 10

1.4 Germplasm Conservation 11Genetic erosion 11Collection and documentation of Mangifera germplasm 12Relevance of germplasm resources to mango improvement 12

1.5 Importance of Mango 12Cultivars 12

1.6 Production and Uses 14

1.1 Introduction

Mango has become a major fruit crop of the tropics and subtropics, particu-larly in Asia, where the mango has always been the most important fruit crop and where it has been considered the king of fruits (Purseglove, 1972). A generation ago, the Green Revolution culminated, creating surpluses of sta-ple and horticultural crops in many developing countries. The Green Revo-lution was the result of nearly a century of effort of applying Mendelian genetics to crop improvement (i.e. conventional breeding) together with the optimization of agronomic and horticultural practices and the successful management of insect pests and diseases. However, improvement of tree

S.K. Mukherjee and R.E. Litz2

crops has lagged far behind eld crops for several reasons: their heterogene-ity, polyploidy, lengthy juvenile period, time required for evaluation of trees in the eld, and the relatively high cost of maintaining tree plantings. For the most part, fruit cultivars continue to be ancient selections, many of which have serious problems, including alternate bearing, lack of disease resistance, low yields, etc. The rapid growth of mango production in recent years has been due to its expansion into new growing regions of the New World, China and parts of Africa; the planting of regular bearing selections; and the adop-tion of modern eld practices, which include irrigation management, control of owering, etc. Agricultural practices are currently undergoing another revolution, as integrated pest and disease management replaces the earlier reliance on agrichemicals, and emerging elds within biotechnology begin to impact cultivar development.

1.2 Description of Mango

The tree

The mango tree is believed to have evolved as a canopy layer or emergent species of the tropical rainforest of South and South-east Asia (Kaur et al., 1980; Bompard, Chapter 2, this volume). Mature trees can attain a height of 40 m or more, and can survive for several hundred years. Mango trees that have been domesticated by selection from openly pollinated seedling popu-lations show variation in tree architecture (i.e. shape and size). The tree is an arborescent evergreen. Leaves are simple and alternate, with petioles that range in length from 1 to 12.5 cm. Leaf morphology is highly variable, de-pending on the cultivar: leaves can be lanceolate, oblong, ovate and interme-diate types involving these forms. Leaf length ranges from 12 to 38 cm and width can be between 213 cm. Young leaves are copper-coloured, changing gradually to light and then dark green with age. The leaves are spirally arranged in whorls and are produced in ushes. The canopy is normally oval, elongated or dome shaped. The juvenile period of seedling trees can range from 3 to 7 years. The root system consists of a long, vigorous taproot and abundant surface feeder roots.

Flowers

Mango owers are borne on terminal pyramidal panicles, and are glabrous or pubescent; the in orescence is rigid and erect, up to 30 cm long, and is widely branched, usually tertiary, although the nal branch is always cymose. The in orescence is usually densely owered with hundreds of small ow-ers, which are 510 mm in diameter. The owers are either monoecious or polygamous, and both monoecious and polygamous owers are borne within a single in orescence (Plate 1). The pistil aborts in male owers. The ratio of monoecious to polygamous owers is strongly in uenced by

Introduction: Botany and Importance 3

environmental and cultural factors. The owers have four or ve sepals and petals that are ovate to ovoid to lanceolate and also thinly pubescent. The oral disc also is four- or ve-lobed, eshy and large and located above the base of the petals. There are ve large, eshy stamens, only one or two of them being fertile; the remaining stamens are sterile staminodes that are sur-mounted by a small gland. In addition, two or three smaller laments arise from the lobes of the nectaries. The stamens are central. The ovule is anatro-pous and pendulous. It is believed that the owers are cross-pollinated by ies (see Davenport, Chapter 5, this volume).

Mukherjee (1951a, 1953) investigated the pollen morphology of mango and 12 other Mangifera species. Their pollen grains were tricolpate of almost the same size. Mondal et al. (1982, cited in Kostermans and Bom-pard, 1993) attempted to correlate pollen morphology with taxonomic relationships of 17 Mangifera species based upon different characteristics of the exine and sporoderm. They demonstrated that all of the species of section II (subgenus Limus) possess coarse exine; whereas there was no clear correlation with pollen type in species within section I (subgenus Mangifera).

The fruit

DescriptionThe mango fruit is a large, eshy drupe, containing an edible mesocarp of varying thickness. The mesocarp is resinous and highly variable with respect to shape, size, colour, presence of bre and avour. The avour ranges from turpentine to sweet. The exocarp is thick and glandular. There is a character-istic beak that develops laterally on the proximal end of the fruit. A sinus is always present above the beak. Fruit shape varies, including elongate, oblong and ovate or intermediate forms involving two of these shapes. Fruit length can range from 2.5 to > 30 cm, depending on the cultivar. The endo-carp is woody, thick and brous; the bres in the mesocarp arise from the endocarp.

The mango fruit is climacteric (see Brecht and Yahia, Chapter 14, this volume), and increased ethylene production occurs during ripening. Chloro-phyll, carotenes, anthocyanins and xanthophylls are all present in the fruit. The skin is generally a mixture of green, red and yellow pigments, although fruit colour at maturity is genotype dependent. During ripening the chloro-plasts in the peel become chromoplasts, which contain yellow and red pig-ments (Krishnamurthy and Subramanyam, 1970; Akamine and Goo, 1973; Salunkhe and Desai, 1984; Mitra and Baldwin, 1997). Peel colour obviously is cultivar dependent (see Knight et al., Chapter 3, this volume). Fruit of Bom-bay Green is green; Carabao, Manila, Mulgoa and Arumanis are greenish-yellow; Dashehari and Alphonso are yellow; and Haden, Keitt and Tommy Atkins have a red blush. The red blush is due to the presence of anthocyanins (Lizada, 1991). The pulp carotenoids in ripe fruit also vary with respect to cultivar (Mitra and Baldwin, 1997).


FlavourFlavour of the mango mesocarp is a function of carbohydrates, organic acids, lactones, monoterpene hydrocarbons and fatty acids (Mitra and Baldwin, 1997). During fruit maturation, starch that accumulates in the chloroplasts is hydrolysed to sucrose, glucose and fructose (Medlicott et al., 1986; Selvaraj et al., 1989; S. Kumar et al., 1994); sucrose is present in slightly higher concen-trations than either fructose or glucose. Organic acid content decreases dur-ing ripening (Krishnamurthy and Subramanyam, 1970). The dominant organic acid is citric acid, but glycolic acid, malic acid, tartaric acid and oxalic acids are also present (Sarker and Muhsi, 1981; Medlicott and Thompson, 1985). The peach-like avour of mangoes is attributed to the presence of lac-tones (Lakshminarayana, 1980; Wilson et al., 1990).

NutritionMango fruit contain amino acids, carbohydrates, fatty acids, minerals, organic acids, proteins and vitamins. During the ripening process, the fruit are ini-tially acidic, astringent and rich in ascorbic acid (vitamin C). Ripe mangoes contain moderate levels of vitamin C, but are fairly rich in provitamin A and vitamins B1 and B2. Perry and Zilva (1932) determined the vitamin A, C and D content of the fruit of three Indian mango cultivars, and found that the pulp of mangoes is a concentrated source of vitamin C. The pulp of mango fruit contains as much vitamin A as butter, although vitamin D is not present in a signi cant quantity. Fruit acidity is primarily due to the presence of malic and citric acids. In addition, oxalic, malonic, succinic, pyruvic, adipic, galac-turonic, glucuronic, tartaric, glycolic and mucic acids are also present (Jain et al., 1959; Fang, 1965). Acidity is cultivar related; for example, immature Florida cultivars have low acidity (0.51.0%) in comparison with Alphonso (3%). During ripening, acidity decreases to 0.10.2%. Following fruit set, starch accumulates in the mesocarp. Free sugars, including glucose, fructose and sucrose, generally increase during ripening; however, the sucrose content increases three- to fourfold due to the hydrolysis of starch. Sucrose is the principal sugar of ripe mangoes. The sucrose content of ripe fruit of three Indian cultivars, Alphonso, Pairie and Totapuri, ranges from 11 to 20% representing 15 to 20% of the total soluble solids (Popenoe, 1932).

The seeds and polyembryony

Mango seeds are solitary, large and at, ovoid oblong and surrounded by the brous endocarp at maturity. The testa and tegumen are thin and papery. Embryos are dicotyledonous. Seeds of monoembryonic mango types contain a single zygotic embryo, whose cotyledons can be unequal in size or lobed in shape. The seeds of polyembryonic mango types contain one or more embryos (Plate 2); usually one embryo is zygotic, whereas the remaining embryos are derived directly from the nucellus, a maternal tissue. Nucellar embryos apparently lack a suspensor. Polyembryony has also been reported in Mangifera casturi, M. laurina and M. odorata (Bompard, 1993). Certain


polyembryonic cultivars reportedly can produce seeds with adventitious nucellar embryos only, for example Strawberry (Juliano, 1934), Carabao and Pico (Juliano and Cuevas, 1932) and Olour and Cambodiana (Maheshwari et al., 1955). Early studies suggested that polyembryony appeared to be a polygenic trait (Juliano, 1934; Sturrock, 1968), segregating as a recessive character in the progeny of controlled crosses. Recent studies, however, have demonstrated that the polyembryony trait is inherited as a dominant character (Aron et al., 1998). Several studies have shown that nucel-lar seedlings can be distinguished from the single zygotic seedling of poly-embryonic seeds by isozymes (Schnell and Knight, 1992; Degani et al., 1993) and DNA markers, for example single sequence repeats (SSRs) (Eiadthong et al., 1999a), ampli ed fragment length polymorphisms (AFLPs) (Kashkush et al., 2001) and inter-simple-sequence-repeats (ISSRs) (Gonzalez et al., 2002). Mango seeds are considered to be recalcitrant, and cannot survive for more than a few days or weeks at ambient temperatures (Parisot, 1988). This important characteristic of mango seeds would have inhibited the long distance dis-persal of mango by seed until recent times.

1.3 History of Cultivation

Origin of Mangifera indica

The largest number of Mangifera species occurs in the Malay Peninsula, the Indonesian archipelago, Thailand, Indochina and the Philippines (Mukher-jee, 1985; Bompard, 1989; see Bompard, Chapter 2, this volume). The most recent classi cation of Mangifera species was based upon oral morphology (Kostermans and Bompard, 1993) and included 69 species, most of which are included in two subgenera Mangifera and Limus with another 11 species occupying an uncertain position (Table 1.1). Eiadthong et al. (1999b) described the phylogenetic relationships among Mangifera species using genomic restriction fragment length polymorphisms (RFLPs) and ampli cation of chloroplast DNA (cpDNA), and suggested that the Mangifera species should be classi ed using molecular data. In the next few years, it is likely that molecular biology will have a major impact on phylogenetic studies involving mango and its relatives.

Mangifera species with a single fertile stamen are distributed in north-eastern India, Myanmar, Thailand and the Malay Peninsula. Many of the mango relatives have small fruits with thin, acidic esh, large seeds, abun-dant bre and astringent resinous substances that are localized near the skin. In addition to M. indica, edible fruit is produced by at least 26 other species in the genus, primarily species found in South-east Asia (Gruezo, 1992). Mangifera caesia, known as binjai or kemang in South-east Asia, is culti-vated in Java, where it bears fruit in the mango off-season (Bompard, 1992a). Mangifera foetida is less commonly cultivated due to its highly astringent fruit; however, the fruit is widely used for pickling and as a substitute for tamarind (Bompard, 1992b). Mangifera kemang and M. altissima are consumed

S.K. M

ukherjee and R.E. Litz

6

Table 1.1. Classi cation of Mangifera species according to Kostermans and Bompard (1993).Genus Subgenus Section Species

Mangifera Mangifera Marchandora Pierre M. gebede MiqEuantherae Pierre M. caloneura Kurz

M. cochinchinensis EnglerM. pentandra Hooker f.

Rawa Kosterm. M. andamanica King M. minutifolia Evard.M. gracilepes M. nicobarica Kosterm.M. grif thii Hooker f. M. paludosa Kosterm.M. merrillii Mukherji M. parvifolia Boerl. & Koorders M. microphylla Griff. ex Hooker f.

Mangifera Ding Hou M. altissima Blanco. M. mucronulata Bl.M. applanata Kosterm. M. oblongifolia Hooker f.M. austro-indica Kosterm. M. orophila Kosterm.M. austro-yunnanensis Hu M. pedicellata Kosterm.M. casturi Kosterm. M. pseudo-indica Kosterm.M. collina Kosterm. M. quadri da JackM. dewildei Kosterm. M. rigida Bl.M. dongnaiensis Pierre M. rubropetala Kosterm.M. ava Evard. M. rufocostata Kosterm.M. indica L. M. similis Bl.M. lalijiwa Kosterm. M. sulauesiana Kosterm.M. laurina Bl. M. sumbawaensis Kosterm.M. linearifolia (Mukherji) Kosterm. M. sylvatica Roxb.M. longipetiolata King M. swintonioides Kosterm.M. magni ca Kochummen M. timorensis Bl.M. minor Bl. M. torquenda Kosterm.M. monandra Merr. M. zeylanica (Bl.) Hooker f.

Introduction: Botany and Im

portance7

Limus (Marchand) Kosterm.

M. blommesteinii Kosterm. M. leschenaultii MarchandM. caesia Jack M. macrocarpa Bl.M. decandra Ding Hou M. odorata Griff.M. foetida Lour. M. pajang Kosterm.M. kemanga Bl. M. superba Hooker f.M. lagenifera Griff.

Species of uncertain position

M. acutigemma Kosterm. M. persiciformis Wu & MingM. bompardii Kosterm. M. subsessifolia Kosterm.M. bullata Kosterm. M. taipa Buch.-HamiltonM. campospermoides Kosterm.

M. transversalis Kosterm.

M. hiemalis Liang Jian Ying M. utana UtanaM. maingayii Hooker f.


as fresh fruit or used green as a salad (Angeles, 1992; Bompard, 1992a). Mangifera pajang has the largest fruit in the genus, and is an attractive fruit. Mangifera odorata is grown in the Philippines and Indonesia, and has occa-sionally been used as a rootstock for mango (Ochse, 1931; Bompard, 1992c). Mangifera odorata is widely grown in the humid lowlands of South-east Asia in areas that are unsuitable for mango as a mango substitute. Mangifera lau-rina and M. pentandra are appreciated as salad ingredients (Bompard, 1992d). In addition, M. grif thii, M. minor, M. monandra, M. quadri da and M. similis have palatable fruit that are considered to have great potential (Gruezo, 1992). All mango cultivars belong to the species M. indica.

According to De Candolle (1884), It is impossible to doubt that it (the mango) is a native of south Asia or of the Malay Archipelago, when we see the multitude of varieties cultivated in those countries, the number of ancient names, in particular a Sanskrit name, its abundance in the gardens of Bengal, of Deccan peninsula, and of Ceylon even in Rheedes time (i.e. 1683). Although the centre of origin and diversity of the genus Mangifera is now rmly established as being in South-east Asia, the origin of M. indica has been a matter of speculation for many years. The fossil record provides few clues, as only a single fossil bearing the imprint of a leaf of M. pentandra has ever been found (Seward, 1912). Mangifera indica is believed to have rst appeared during the Quatenary period (Mukherjee, 1951b). Blume (1850) considered that mango might have originated from several related species, primarily located in the Malay archipelago.

On the basis of ancient accounts of travellers and the written historical record, it was believed for many years that mango must have originated in India and spread outwards from there to South-east Asia and thence to the New World and Africa. Because north-eastern India is at the northernmost edge of the distribution of the Mangifera species, Hooker (1876) suggested that mango might have been naturalized in India. The historical record pro-vides a sometimes con icting account of the distribution of mango. Miquel (1859) did not record it as being wild in the Indonesian archipelago. Accord-ing to Rumphius (1741), the mango was introduced into certain islands of the Indonesian archipelago within recent times; however, the mango was in cul-tivation in Java at least as early as ad 9001100, when the temple at Borobo-dur was built and faced with carvings of the Buddha in contemplation under a mango tree (Plate 3). Based upon taxonomic and recent molecular evidence, it is now apparent that the mango probably evolved within a large area including north-western Myanmar, Bangladesh and north-eastern India (see Bompard, Chapter 2, this volume).

Polyembryonic and monoembryonic M. indicaWithin M. indica, there are two distinct types that can be distinguished on the basis of their mode of reproduction and their respective centres of diversity: a subtropical group with monoembryonic seed (Indian type) and a tropical group with polyembryonic seed (South-east Asian). A few polyembryonic cultivars occur along the west coast of India; however, they may have been introduced into Goa from South-east Asia, perhaps by the Portuguese from


their colonies of Malacca in the Malay Peninsula or Timor in the Indonesian archipelago. Kumar et al. (2001) estimated the genetic relatedness among ten polyembryonic and monoembryonic cultivars from the west coast of south-ern India using genomic and chloroplast DNA RFLP analysis. The cultivars could be grouped on the basis of embryo type (i.e. monoembryonic and poly-embryonic) and had distinctly different genetic backgrounds. They con-cluded that polyembryonic mangoes could not have originated in India, and must have been introduced, probably from South-east Asia.

Domestication of mango

Historical recordIt is probable that mango cultivation originated in India, where De Candolle (1884) estimated that mango cultivation appeared to have begun at least 4000 years ago. In the early period of domestication, mango trees probably yielded small fruit with thin esh. Such fruit can be found today in north-eastern India and in the Andaman Islands (Anonymous, 1992). Folk selections of superior seedlings over many hundreds of years would have resulted in larger fruit with thicker esh. Mukherjee (1950a, b) described many of these primitive selections from Orissa in north-eastern India; they demonstrated great variation in fruit shape and size.

The mango is a very important cultural and religious symbol of India. Buddhist pilgrims Fa-Hien and Sung-Yun mentioned in their travel notes that the Gautama Buddha was presented with a mango grove by Amradarika (c.500 bc) as a place for meditation (Popenoe, 1932). According to Burns and Prayag (1921), a mango tree is depicted in friezes on the stupa of Bharut, which was constructed c.100 bc. Other travellers to India, including the Chi-nese Hwen Tsung (ad 632645), the Arabs Ibn Hankal (ad 902968) and Ibn Batuta (ad 13251349) and the Portuguese Lurdovei de Varthema (ad 15031508), all described the mango. The Indian subcontinent was the birthplace of some of the earliest highly developed civilizations, and over the centuries, India exerted strong cultural, religious and commercial in uence over South and South-east Asia. In successive waves, Hinduism, Buddhism and Islam were introduced into South-east Asia from India. To this day, many com-monly used words in Indonesia are derived from both Sanskrit and Tamil. One of the most widely used words for mango in Malaysia and Java (Indone-sia) is mangga, which is derived from the Tamil manga. Traders and monks from India possibly introduced superior selections of mango into South-east Asia; however, vegetative propagation was unknown in India until after the arrival of the Portuguese in Goa in the 15th century. Moreover, the most im-portant mango selections of Thailand, Cambodia, Vietnam, Malaysia, Indo-nesia and the Philippines historically have all been of the polyembryonic type, and have traditionally been seed propagated. Until the establishment of Portuguese enclaves on the coast of India beginning in the late 15th century, mango cultivars did not exist in India, as there was no known method for vegetatively propagating superior selections (see Iyer and Schnell,


Chapter 4, this volume). However, under the Moghul emperor Akbar (15561605), the best selections of seedling mangoes were propagated by approach grafting and were planted in large orchards. The Lakh Bagh, a mango orchard of 100,000 trees, was planted near Darbhanga in Bihar. Perhaps noth-ing more eloquently attests to the importance of this fruit and the esteem in which it was held than this vast mango orchard. The Ain-i-Akbari, an ency-clopedic work that was written during the reign of Akbar, contains a lengthy account of the mango, and includes information about the quality of the fruit and varietal characteristics. There was evidently a strong body of informa-tion about mango cultivation that had accumulated up to that time. Most of the mango cultivars of India had their origin in those years, and have been maintained under cultivation for over 400 years by vegetative propagation. Alphonso, Dashehari, Langra, Rani Pasand, Safdar Pasand and other mango cultivars were selected during that time. Relics of orchards from the time of Akbar are found in different parts of India, and it has been suggested that they could still provide valuable material for selection of superior mango cultivars.

Distribution

Spreading from the centres of domesticationThe global spread of mangoes and their cultivation outside their original centres of domestication probably did not occur until the beginning of the European voyages of discovery and colonialization in the 15th and 16th centuries. Because mango seeds are recalcitrant, and cannot survive for more than a few days or weeks, mango germplasm in the early days must have been transported as ripe fruit, seedlings or, later on, as grafted plants. It is believed that the Portuguese transported the mango from their colonies in India to their African colonies, although Purseglove (1972) suggested that it might also have been introduced to Africa via Persia and Arabia in the 10th century by Arab traders. The Portuguese later introduced the mango into Brazil from their African colonies of Mozambique and Angola. Spaniards, who encountered a mango-growing civilization in the Philippines after Magellans passage across the Paci c Ocean, introduced polyembryonic mango types to their New World colonies through the Paci c trading ports of Mexico and Panama. The most important, traditional mango cultivar in Mexico remains the Manila, re ecting its Philippine origin. Carabao and Manila are probably identical. The mango was introduced to the West Indies in the mid- to late 18th century, probably from Brazil. The rst introductions of mango into Florida (USA) occurred in 1861, and involved the No. 11 polyembryonic seedling from Cuba. Seven years later, another polyembry-onic selection, Peach was introduced into the state (Knight and Schnell, 1993). Many of the early introductions into Florida proved to be unproduc-tive, although Mulgoba was planted on a small commercial scale (this culti-var is referred to as Mulgoa in India, Mulgoba in the USA and Malgoa in Malaysia).


Secondary centres of diversityIn 1910, a seedling of Mulgoba came into production in Florida. Its fruit had a highly attractive red blush, and appeared to bear more heavily than its parent(s) (Wolfe, 1962). This selection was named Haden. Although Haden was not superior with respect to fruit quality in comparison to the imported germplasm from India, its genetic base was much wider. During the 20th century, more introductions of mango germplasm into Florida occurred from South-east Asia (the Philippines, Cambodia), India and elsewhere. It was at one time believed that these introductions of mango germplasm created a secondary centre of diversity of the species (Knight and Schnell, 1993). Eldon, Glenn, Lippens, Osteen, Parvin, Smith, Springfels, Tommy Atkins and Zill are progeny of Haden. Saigon seedlings were selections made from Cambodiana, a polyembryonic introduction from Indochina. From Saigon seedlings, Alice, Herman and Florigon were selected. Based upon more recent genetic analysis involving microsatellite markers, it is now estimated that the majority of Florida cultivars are descended from only four monoembryonic Indian mango cultivar accessions, i.e., Mulgoba, Sandersha, Amini and Bombay, together with the polyembryonic Tur-pentine from the West Indies (Schnell et al., 2006). The Florida mango culti-vars have been found to be highly adaptable to many agroecological areas and bear regularly, whereas many of the outstanding Indian cultivars have been unproductive outside their centre of domestication, and are alternate bearing. These selections also have a highly attractive red blush at maturity, rm esh, a high esh to seed ratio and a regular bearing habit. Some of the Florida cultivars, for example Tommy Atkins, Keitt, etc. are also moder-ately resistant to anthracnose, the most important production and posthar-vest problem of mango in many areas. In the latter half of the 20th century, plantings of Florida cultivars have been established in many countries and now form the basis of international trade of mangoes.

Current distributionThe mango is cultivated commercially throughout the tropics and in many subtropical areas. It is grown at the equator and at a latitude of 3537q in southern Spain. According to Knight and Schnell (1993), The process that began in Florida introduction of superior germplasm from abroad followed by selection of improved cultivars adapted to local conditions is now underway in many areas.

1.4 Germplasm Conservation

Genetic erosion

The Mangifera species have their centre of diversity and origin in South-east Asia, a region that has experienced great economic development in recent years. Vast wooded areas have been completely or partially deforested either for expanding agriculture or for removal of tropical hardwoods for export.


This has caused great genetic erosion within many species and genera. The Mangifera species, like many other tropical fruit trees, are canopy and emer-gent trees of the tropical rainforest (Kaur et al., 1980). These trees are widely scattered in the tropical rainforest, ower erratically and reproduce from large seeds that deteriorate rapidly. As such, they are particularly vulnerable and in danger of extinction.

Collection and documentation of Mangifera germplasm

The International Plant Genetic Resources Institute (IPGRI), formerly known as the International Board for Plant Genetic Resources (IBPGR), commis-sioned an ecogeographical study of known Mangifera genetic resources (Muk-herjee, 1985). Based upon this documentation, a joint IBPGR-International Union for the Conservation of Nature (IUCN)-World Wildlife Fund (WWF) project was initiated to collect wild mangoes on the island of Borneo and in the Malay Peninsula (Bompard, 1989), the regions that held the highest con-centrations of Mangifera species. Kostermans and Bompard (1993), in the lat-est revision of the taxonomy of Mangifera, recognized 69 species, many of which were collected during the course of this project (Table 1.1). Because of the loss of natural habitat, the establishment of in situ and ex situ germplasm collections of Mangifera species was considered to be imperative.

Relevance of germplasm resources to mango improvement

The genetic improvement of mango hitherto has depended on the utilization of the genetic variability found within a single species, M. indica. According to Mukherjee (1985), A concerted sampling strategy should be devised for exsitu samples to meet urgent needs for use in research for improvement of the crop through breeding or as rootstocks. Sources of resistance to mango mal-formation, anthracnose, powdery mildew, gall midge are urgently needed.

1.5 Importance of Mango

Cultivars

A partial list of the principal mango cultivars has been provided in Table 1.2. This list includes many cultivars that were identi ed in a survey of world mango production compiled by Watson and Winston (1984). The distribution of mango cultivars outside their centres of domestication can be attributed primarily to three historical events: (i) the movement of Indian varieties (monoembryonic) along the trade routes of the Portuguese to Africa and South America; (ii) the spread of South-east Asian varieties (polyembryonic) across the Paci c Ocean to Central and South America by the Spaniards; and (iii) the identi cation of improved mango cultivars initially in Florida and


Table 1.2. Most important mango cultivars in major producing countries.

Continent Country Cultivars

Africa Cote dIvoire Amelie, KentEgypt Alphonso, Bullocks Heart, Hindi be Sennara,

Langra, Mabrouka, Pairie, Taimour, ZebdaKenya Boubo, Ngowe, BatawiMali Amelie, KentSouth Africa Fascell, Haden, Keitt, Kent, Sensation, Tommy

Atkins, ZillAsia Bangladesh Aswina, Fazli, Gopal Bhog, Himsagar, Khirsapati,

LangraChina Gui Fei, Tainong No. 1, Keitt, Sensation, Zill, Zihua,

Jin HuangIndia Alphonso, Banganapalli, Bombay, Bombay Green,

Chausa, Dashehari, Fazli, Fernandian, Himsagar, Kesar, Kishen Bhog, Langra, Mallika, Mankurad, Mulgoa, Neelum, Pairi, Samar Behisht, Suvarnarekha, Totapuri, Vanraj, Zardalu

Indonesia Arumanis, Dodol, Gedong, Golek, 'Madu, 'ManalagiIsrael Haden, Tommy Atkins, Keitt, Maya, Nimrod, Kent,

PalmerMalaysia Apple Rumani, Arumanis, Golek, Kuala Selangor 2,

MalgoaMyanmar Aug Din, Ma Chit Su, Sein Ta Lone, 'Shwe Hin ThaPakistan Anwar Ratol, Began Pali, Chausa, Dashehari,

Gulab Khas, Langra, Siroli, Sindhri, Suvarnarekha, Zafran

The Philippines Carabao, Manila Super, PicoTaiwan Irwin, Jin-hwung, Keitt, Tommy Atkins, Tainong

No. 1, Tsar-swainThailand Nam Doc Mai, Ngar Charn, Ok Rong, Keow Savoey,

Pimsen MumAustralia Calypso, Kensington PrideNorth and

Central America

Costa Rica Haden, Irwin, Keitt, Mora, Tommy AtkinsDominican Republic

Haden, Keitt, Kent, Tommy Atkins

Guatemala Haden, Keitt, Kent, Tommy AtkinsHaiti Francine, Madame FrancisMexico Ataulfo, Haden, Keitt, Kent, Manila, Palmer,

Sensation, Tommy Atkins, Van DykeUSA Keitt, Kent, Tommy Atkins

SouthAmerica

Brazil Bourbon, Coite, Coquinho, Coracao, Espada, Haden, Itamaraca, Keitt, Mamao, Palmer, Rosa, Tommy Atkins, Uba, Van Dyke

Colombia VallenatoEcuador Haden, Keitt, Kent, Tommy AtkinsPeru Haden, Keitt, Kent, Tommy AtkinsVenezuela Haden, Keitt, Kent, Tommy Atkins


later in other new mango-producing areas, as a result of open and controlled pollination among local and introduced mango germplasm from India and South-east Asia.

Further information about many of the mango cultivars, including their fruit characters, is available in Knight et al. (Chapter 3, this volume), and in publications by Burns and Prayag (1921) for mangoes of Maharashtra, Naik and Gangolly (1950) for south Indian mangoes, Singh and Singh (1956) for Uttar Pradesh mangoes, Mukherjee (1948) for Bengal mangoes and Camp-bell (1992) for Florida mangoes.

Because many clonally propagated mango cultivars have unique local and/or regional names, there is considerable confusion in nomenclature. The Indian Agricultural Research Institute (IARI), New Delhi, has been recog-nized by the International Society for Horticultural Science (ISHS) as the International Registration Authority for Mango, whose mission is to consoli-date super uous names of mango cultivars. The potential for molecular, for example randomly ampli ed polymorphic DNA (RAPD), markers, to resolve much of this confusion has been demonstrated by Schnell and Knight (1992), Degani et al. (1993), Schnell et al. (1995), Eiadthong et al. (1999a), Kashkush et al. (2001) and Gonzalez et al. (2002) (see Bompard, Chapter 2 and Iyer and Schnell, Chapter 4, this volume).

There is little variation among seedlings derived from polyembryonic mangoes. None the less, a certain amount of variability does occur, probably as a result of somatic mutation. Thus, in Indonesia there are several Aru-manis selections that are denoted numerically, for example Arumanis 1, Arumanis 2, etc. In addition, although Philippine mango cultivars are dis-tinguished by different names, for example Carabao, Manila, Philippine, etc., the differences among them are quite subtle.

1.6 Production and Uses

The mango is the most important fruit of Asia, and currently ranks fth in total production (in metric tonnes) among major fruit crops worldwide, after Musa (bananas and plantains) (105,815,354 t), Citrus (all types) (105,440,168 t), grapes (65,584,233 t) and apples (59,444,377 t) (FAOSTAT, 2006). According to the Food and Agriculture Organization of the United Nations (FAO) database (FAOSTAT, 2006), world mango production has increased from 16,903,407 t in 1990 to 28,221,510 t in 2005. Much of this new production has occurred outside the traditional centres of mango culture of South and South-east Asia. In 1990, India produced approximately 51% of the worlds mangoes, but by 2005, Indias share had declined to approximately 38%, despite the substantial increase in mango production since 1990 (from 8,645,405 to 10,800,000 t between 1990 and 2005). The current leading producing nations after India include (in metric tonnes) China (3,450,000), Thailand (1,800,000), Pakistan (1,673,900), Mexico (1,600,000), Indonesia (1,478,204), Brazil (1,000,000) and the Philippines (950,000). Although world production has increased by 67% between 1990 and 2005, mango exports have increased almost sixfold


from 158,030 to 907,782 t, with total export value estimated to be US$583,763,000 (FAOSTAT, 2006). The major exporting countries are (in met-ric tonnes) Mexico (212,505), India (156,222) and Brazil (111,181). As a result, mangoes are widely available as fresh fruit and as processed products (i.e. dried fruit, dairy products, juice, pickles, etc.).

Mangoes are an important component of the diet in many less developed countries in the subtropics and tropics. In regions of the world that have experienced low living standards and serious nutritional de ciencies, their attractiveness and avour have also enhanced the quality of life. Surplus production has increasingly been processed and fruit of certain cultivars is destined for export as fresh fruit. Approximately 1% of mango production is utilized for processing for juice, nectars, preserves (including chutney), fruit leather, dried fruit slices, frozen pulp and as a avouring for baked goods, ice cream, yoghurt, etc. (see Raymundo et al., Chapter 17, this volume). No part of the fruit is wasted. In India and the subcontinent, the seed is used for extraction of starch amchur, and the peels (skin) have been used as a source of anacardic acid. Mango wood is a low quality timber, and the bark of the tree is an important source of tannins for curing leather.

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characteristics of zygotic seedlings from polyembryonic mango cultivars, deter-mined using isozymes as genetic markers. Acta Horticulturae 341, 7885.

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Eiadthong, W., Yonemoni, K., Sugiura, A., Utsunomiya, N. and Subhadrabandhu, S. (1999b) Analysis of phylogenetic relationship in Mangifera by restriction site analy-sis of an ampli ed region of cpDNA. Scientia Horticulturae 80, 145155.

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Hooker, J.D. (1876) The Flora of British India 2. Reeve, London.Jain, N.L., Krishnamurthy, G.V. and Lal, G. (1959) Nonvolatile organic acids in unripe

pickling mangoes and salted mango slices by paper chromatography. Food Science8, 115117.

Juliano, J.B. (1934) Origin of embryos in the strawberry mango. Philippine Journal of Science 54, 553563.

Juliano, J.B. and Cuevas, N.L. (1932) Floral morphology of the mango (Mangifera indicaL.) with special reference to the Pico variety from the Philippines. Philippine Agri-culturist 21, 449472.

Kashkush, K., Jinggui, F., Tomer, E., Hillel, J. and Lavi, U. (2001) Cultivar identi cation and genetic map of mango (Mangifera indica). Euphytica 122, 129136.

Kaur, A., Ha, C.O., Jong, K., Sands, V.E., Chan, H.T., Soepadmo, E. and Ashton, P.S. (1980) Apomixis may be widespread among trees of the climax rain forest. Nature271, 440442.

Knight, R.J., Jr and Schnell, R.A. (1993) Mango (Mangifera indica L.) introduction and evaluation in Florida and its impact on the world industry. Acta Horticulturae 341, 125135.

Kostermans, A.J.G.H. and Bompard, J.M. (1993) The Mangoes: Botany, Nomenclature, Horticulture, Cultivation and Utilization. Academic Press, London.

Krishnamurthy, S. and Subramanyam, H. (1970) Respiratory climacteric and chemical changes in the mango fruit Mangifera indica L. Journal of the American Society for Horticultural Science 95, 333337.

Kumar, N.V.H., Narayanaswamy, P., Prasod, D.T., Mukunda, G.K. and Sondhu, S.N. (2001) Estimation of genetic diversity of commercial mango (Mangifera indica L.)cultivars using RAPD markers. Journal of Horticultural Science and Biotechnology 76, 529533.


Kumar, S., Das, D.K., Singh, A.K. and Prasad, U.S. (1994) Sucrose metabolism during maturation and ripening of mango cultivars. Plant Physiology and Biochemistry 21, 2732.

Lakshminarayana, S. (1980) Mango. In: Nagy, S. and Shaw, P.E. (eds) Tropical and Subtropical Fruits. AVI Publishing Co., Westport, Connecticut, USA, pp. 184257.

Lizada, M.C. (1991) Post harvest physiology of mango a review. Acta Horticulturae 291, 437449.

Maheshwari, P., Sachar, R.C. and Chopra, R.N. (1955) Embryological studies in mango (Mangifera indica L.). In: Proceedings of the 42nd Indian Scienti c Congress, Baro-da, India, p. 233.

Medlicott, A.P. and Thompson, A.K. (1985) Analysis of sugars and organic acids in ripening mango fruit (Mangifera indica L. var. Keitt) by high performance liquid chromatography. Journal of the Science of Food and Agriculture 36, 56566.

Medlicott, A.P., Bhogol, M. and Reynolds, S.B. (1986) Changes in peel pigmentation during ripening of mango fruit (Mangifera indica var. Tommy Atkins). Annals of Applied Biology 109, 651656.

Miquel, F.A.G. (1859) Flora van Nederlandsch Indie 1, 627634.Mitra, S.K. and Baldwin, E.A. (1997) Mango. In: Mitra, S.K. (ed.) Postharvest Physiology

and Storage of Tropical and Subtropical Fruits. CAB International, Wallingford, UK, pp. 85122.

Mukherjee, S.K. (1948) The varieties of mango (M. indica L.) and their classi cation. Bulletin of the Botanical Society of Bengal 2, 101133.

Mukherjee, S.K. (1950a) Wild mangoes of India. Science and Culture 15, 469471.Mukherjee, S.K. (1950b) Mango. Its allopolyploid nature. Nature 150, 196197.Mukherjee, S.K. (1951a) Pollen analysis in Mangifera in relation to fruit set and taxonomy.

Journal of the Indian Botanical Society 30, 4955.Mukherjee, S.K. (1951b) Origin of mango. Indian Journal of Genetics and Plant Breeding

11, 4956.Mukherjee, S.K. (1953) Origin, distribution and phylogenetic af nities of the species of

Mangifera L. Journal of the Linnean Society, Botany 55, 6583.Naik, K.C. and Gangolly, S.R. (1950) A Monograph on Classi cation and Nomenclature

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(Jakarta), Indonesia.Parisot, E. (1988) Etude de la croissance rhythmique chez de jeunes manguiers (Mangifera

indica L.). Description, germination et conservation de graines polyembryonnees de manguier. Fruits 43, 97105.

Perry, E.O.V. and Zilva, S.S. (1932) Preliminary Report on Vitamin Content of the Mango.Empire Marketing Board, London.

Popenoe, W. (1932) Manual of Tropical and Subtropical Fruits. Macmillan Co., New York.

Purseglove, J.W. (1972) Mangoes west of India. Acta Horticulturae 24, 107174.Rumphius, G.E. (17411750) Herbarium Amboinense. Vol. 16. Den Haag, Amsterdam.Salunkhe, D.K. and Desai, B.B. (1984) Mango. In: Postharvest Biotechnology of Fruits,

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ganic acids of mango (Mangifera indica L.) at various stages of fruit development. Bangladesh Journal of Agricultural Science 8, 6975.

Schnell, R.J. and Knight, R.J., Jr (1992) Frequency of zygotic seedlings from ve polyem-bryonic mango rootstocks. HortScience 27, 174176.


Schnell, R.J., Ronning, C.M. and Knight, R.J., Jr (1995) Identi cation of cultivars and validation of genetic relationships in Mangifera indica L. using RAPD markers. Theoretical and Applied Genetics 90, 269274.

Schnell, R.J., Brown, J.S., Olano, C.T., Meerow, A.W., Campbell, R.J. and Kuhn, D.N. (2006) Mango genetic diversity analysis and pedigree inferences for Florida culti-vars using microsatellite markers. Journal of the American Society for Horticultural Science 13, 214224.

Selvaraj, Y., Kumar, R. and Pal, D.K. (1989) Changes in sugars, organics, amino acids, lipids, lipid constituents and aroma characteristics or ripening mango (Mangiferaindica L.) fruit. Journal of Food Science and Technology 26, 306311.

Seward, A.C. (1912) Dictyledonous leaves from Assam. Records of the Geological Sur-vey of India 42, 100.

Singh, L.B. and Singh, R.N. (1956) A Monograph on the Mangoes of UP. Superintendent of Printing, Uttar Pradesh Government, Lucknow, India.

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CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses(ed. R.E. Litz) 19

2 Taxonomy and Systematics

J.M. BompardLes Mazes, Montaud, France

2.1 Introduction 192.2 The Genus Mangifera L. 20

Distribution 20Ecology and habitat 20

2.3 Taxonomy and Systematics 22Taxonomic history 22

2.4 Phytogeography 28Species distribution 28Subgenera and section distribution 29

2.5 Interspeci c Molecular Characterization 302.6 Region of Origin of the Genus 312.7 Origin of the Common Mango 32

The common mango in South-east Asia 322.8 Conclusion 35

Potential contribution of wild species to mango cultivation 35Source of rootstock 35Hybridization 36Potential of wild species 36

2.1 Introduction

The genus Mangifera is one of the 73 genera (c.850 species) belonging to the family of Anacardiaceae, in the order of Sapindales. Anacardiaceae is a fam-ily of mainly tropical species, with a few representatives in temperate regions. Malesia, which is the phytogeographic region extending from the Malay Peninsula south of the Kangar-Pattani line to the Bismarck Archipelago east of New Guinea (Whitmore, 1975) contains more species in the Anacardiaceae than any other area. Within Malesia occurrence is mainly in Western Malesia (Ding Hou, 1978b).

J.M. Bompard20

Apart from mango, several other cultivated fruit trees belong to the fam-ily, for example the ambarella or Otaheite apple (Spondias dulcis Forst.) prob-ably from Melanesia, and the yellow and purple mombins (Spondias mombin L. and S. purpurea L., respectively) from tropical America, the Bouea species from IndoMalesia, dragon plums (Dracontomelum spp.) from IndoMalesia and the Paci c region, kaf r plum (Harpephyllum caffrum Bernh. ex K. Krause) and the marula plum (Sclerocarya caffra Sond.) of southern Africa. The cashew (Anacardium occidentale L.) is from tropical America and the pistachio (Pistaciavera L.) from Iran and Central Asia. Anacardiaceous species also yield other valuable products: wood (several genera), gums and resins (Pistacia spp.), varnishes (Rhus spp. and Melanorrhoea spp., lacquer trees) and tanning materials (Rhus spp. and Schinopsis spp.). It is also a family well known for the dermal irritation produced by some of its members, such as the poison ivies and oaks (Rhus spp.) in North America, rengas (Gluta spp.) in South-east Asia and other species including some Mangifera species whose resinous sap may induce a mild to strong allergic reaction.

2.2 The Genus Mangifera L.

Distribution

The range of natural distribution of the 69 Mangifera species is mainly restricted to tropical Asia, and extends as far north as 27 latitude and as far east as the Carolines Islands. Wild mangoes occur in India, Sri Lanka, Ban-gladesh, Myanmar, Sikkim, Thailand, Kampuchea, Vietnam, Laos, southern China, Malaysia, Singapore, Indonesia, Brunei, the Philippines, Papua New Guinea and the Solomon and Carolines Islands. The highest species diver-sity, c.29 species, occurs in western Malesia, especially in peninsular Malay-sia and in Borneo and Sumatra, which represent the heart of the distribution range of the genus (Fig. 2.1).

Ecology and habitat

The majority of Mangifera species occur as a rule as scattered individuals in tropical lowland rainforests on well-drained soils. The species are distrib-uted mostly below 300 m, but can occur up to c.1000 m above sea level, on well-drained soils (44 species), in periodically inundated areas (ten species) and in certain types of swamp forest (i.e. M. gedebe, M. grif thii and M. parvi-folia). Three species are mainly found in sub-montane forests above 1000 m and occasionally up to 1700 m above sea level (M. bompardii, M. dongnaiensis and M. orophila). There are also species that are adapted to seasonally dry climates in deciduous or semi-deciduous forests (e.g. M. caloneura, M. collina, M. timorensis and M. zeylanica). A few species occur north of the Tropic of Cancer, for example M. austro-yunnanensis and M. persiciformis in China, M.

Taxonomy and Systematics 21

sylvatica Roxb. in Sikkim and southern China, at altitudes of 6001900 m above sea level; apparently wild M. indica can also be found outside the tropics.

Wild mangoes are large trees, 3040 m (occasionally 54 m) in height, with tall columnar boles. Several species are exploited for their timber. The major-ity of wild mangoes occur as scattered individuals at very low densities in lowland forests on well-drained soils. Some of these are very rare; there are normally one to three trees above 40 cm in diameter/10 ha. Only a few spe-cies (M. gedebe, M. grif thii and M. parvifolia) are gregarious in certain types of swamp forest. Most species are evergreen although a few are deciduous in the rainforests following a dry period, and stand bare for a short time before ushes of new leaves appear. A deciduous habit that is not linked to a sea-sonal climate also occurs in other genera of Anacardiaceae (Ding Hou, 1978b).

In the rainforest of western Malesia, Mangifera species ower and fruit very irregularly. As with many other genera in the region, mast or general fruiting at intervals of 38 years is the dominant pattern. In mast years, the ground beneath the trees can be covered with mangoes, whose strong smell attracts many animals. Isolated owering may occur after a dry period and is generally followed by a poor fruit crop. The occurrence of owering of a few species, for example the lanjut (M. lagenifera) is only once every 510 years. There seem to be clear reproductive barriers between species in the wild, although limited hybridization among cultivated species has been reported (see section 2.8, Conclusion, this chapter).

30 N

4

2

5

4

3

613

27

2728

95

7 4

3

5

2

20 N

10 N

0 N

10 N

30 N

20 N

10 N

0 N

10 N

80 100 120 160

80 100 120 160140

Fig. 2.1. Distribution of Mangifera species in the range of the genus. Numbers shown indicate the number of wild species in each area: Sri Lanka, India and Sikkim, Andaman and Nicobar Islands, Myanmar, Thailand, Indochina, China, peninsular Malaysia, Sumatra, Borneo, Java, Lesser Sunda Islands, Sulawesi, Moluccas, the Philippines, New Guinea, and Solomon Islands (the Caroline Islands not represented).

J.M. Bompard22

These widely scattered towering tree species, often with an inaccessible crown, are undercollected and poorly represented in herbarium collections (Bompard, 1995). Because of their irregular owering, the owers and fruits of a few species are still unknown. Collecting plant material is consequently very dif cult, and plant explorations are still yielding new records or new species. Many species have been recently recorded for the rst time, even from peninsular Malaysia, a country that has already been rather well combed by botanists, having one of the highest collecting indices in the Malesian region. Other species still await to be discovered. Sadly some species of very limited range may already have been lost to posterity by deforestation.

Our very meagre knowledge of the wild mangoes is due to the fact that identi cation at the species level from leaves only is often dif cult because of intraspeci c variation in vegetative characters. Moreover, many of the origi-nal species were based on very poor specimens. Consequently, frequent mis-identi cation of herbarium material has resulted in much confusion, requiring a critical revision of all the specimens in these collections. It is not uncommon that the same species has been described from different places under differ-ent names. For instance, M. inocarpoides described by Merrill and Perry from New Guinea in 1941, M. camptosperma and M. reba (recorded by Pierre in South Vietnam in 1897) are now recognized to be a single species M. gedebe Miquel, a species initially named in 1861 from a specimen collected in Suma-tra. Mangifera longipes Grif th is now treated as M. laurina Blume, because this name takes precedence as it was validly published 4 years earlier.

After thorough study of herbarium collections and eld collections, a number of species have been newly described. Sixty-nine species are now recorded, including 13 species of uncertain af nities, in contrast with the 49 species recognized by Mukherjee (1949). As more collections are made, there will doubtless be further taxonomic adjustments made to the genus Mangifera.

2.3 Taxonomy and Systematics

Taxonomic history

Subdivision of the genusAn historical review of the subdivisions of the genus Mangifera shows that two major groups have been rather consistently recognized in taxonomic treatments. Hooker (1862) was the rst to recognize two sections based on the characters of the ower disc: section I with a disc broader than the ovary, and section II with a disc stalk-like or wanting. These sections were later named by Marchand (1869) Amba, an Indian name for the common mango, and Limus, a Sundanese name for M. foetida in West Java, respectively. He also added a section Manga for M. leschenaultii, which in fact belongs to the section Limus.

In his monograph of the Anacardiaceae, Engler (1883) maintained Hook-ers sections, and subdivided group A (Hookers section I) into two groups,


one group with four or ve petals and the other group with four petals. He considered the following sequence of morphological characters to be impor-tant for classi cation: (i) texture of the leaves; (ii) number of fertile stamens; (iii) prominence of veins; (iv) pilosity of in orescences; and (v) leaf shape.

Pierre (1897) further divided the genus Mangifera into ve sections based on ower characters, i.e. number of stamens, the attachment of stamens to the disk, and the style. Two of these ve sections namely section I Euan-therae, with a short thick ower disc and 412 fertile stamens, and section V Marchandora then consisting of M. camptosperma (currently considered a syn-onym of M. gedebe) are still maintained as they form clear-cut sections.

In his monograph, Mukherjee (1949) recognized two unnamed sections, conserving Hookers subdivision. Ding Hou (1978a) adopted the same method in his revision of the Malesian Anacardiaceae recognizing only Hookers two original sections and providing them with proper names and synonyms: section Mangifera (section I Hooker, section Amba Marchand, group A Engler, sections Euantherae and Marchandora Pierre) and section Limus (section II Hooker, sections Limus and Manga Marchand, group B Engler, and sections Eudiscus and Microdiscus Pierre).

Most recent classi cation of the genusThe taxonomic classi cation referred to herein follows that proposed by Kos-termans and Bompard (1993). This treatise includes the results of collections and surveys carried out between 1986 to 1998 in Borneo and peninsular Malaysia, which were initiated and sponsored by the International Institute for Genetic Resources (now Biodiversity International) and the World Wide Fund for Nature.1 It was published under the auspices of the International Board for Plant Genetic Resources (now International Plant Genetics Resources Institute) and the Linnean Society of London.

The most recent treatment of Mangifera re ects the current status of what is still fragmentary knowledge. It can provide a basis for further studies involving all aspects of the wild relatives of mango, but particularly their potential in mango breeding. Determining phylogenetic af nities based upon molecular markers could change our thinking about relationships among Mangifera species and among the cultivated forms of M. indica (see Interspe-ci c Molecular Characterization section, this chapter).

The morphological characters used for identi cation have been placed in the following sequence of importance:

Shape of the oral disc (see section Subdivision of the genus).1. Number of fertile stamens.2. Seed labyrinthine or not.3. Shape of secondary branches of the in orescences: open or lax panicle, 4.

owers glomerulate or sub-glomerulate, the rami cations racemoid or spike like.

Pubescence of the in orescence.5. Shape, number and attachment of the nerves (ridges or ngers) at the 6.

inner surface of the petals.

J.M. Bompard24

Shape and size of the petals.7. Flowers tetra- or pentamerous (not a very constant character and often 8.

overlapping).Reticulation of the leaves, especially of the lower surface.9. Shape of the leaf (only fully grown leaves of sterile branches can be used).10. Texture of the leaves.11. Deciduous or non-deciduous trees.12. Colour of the owers.13. Shape, colour and smoothness of the fruit.14. Number and size of the stone bres.15.

Kostermans (Kostermans and Bompard, 1993) raised the sections to the rank of subgenus, i.e. subgenus Limus (Marchand) Kosterm., having a disc narrower than the base of the ovary, stalk-like or even lacking and subgenus Mangifera (Ding Hou) Kosterm., having a disc broader than the base of the ovary, cushion-like, often divided in four or ve lobes.

SUBGENUS LIMUS (MARCHAND) KOSTERM. Mangifera species of the subgenus Limus are quite distinctive and show only remote af nity with the common mango. This taxon is more primitive than the subgenus Mangifera and may be ances-tral to it, although the two subgenera may have originated from two different ancestors. The subgenus Limus consists of 11 species, which are native to the rainforests of western Malesia (peninsular Thailand, Malay Peninsula, Suma-tra, West Java and Borneo), with the exception of M. foetida, which extends to the east, possibly as far as New Guinea, and M. odorata which is only known in cultivation.

Kostermans divided the subgenus Limus into two sections: (i) section Deciduae for deciduous trees (i.e. M. caesia, M. kemanga, M. pajang, M. superba and possibly M. blommesteinii, M. decandra and M. lagenifera); and (ii) section Perennes for non-deciduous species (i.e. M. foetida, M. leschenaultii, M. macro-carpa and M. odorata) (Kostermans and Bompard, 1993). In deciduous trees, the bracts enclosing the buds leave a characteristic collar of dense, narrow scars, which persist on old twigs and are especially prominent in M. caesia and M. kemanga.

Mangifera lagenifera and M. decandra have ten stamens, ve of which are fertile. The other nine species have only one (and rarely two) fertile stamen(s) and two to four staminodes. The two species with ve fertile stamens (M.decandra and M. lagenifera) and M. superba, M. caesia, M. kemanga and M. blom-mesteinii, whose leaves are apically aggregated into rosettes at the end of mas-sive twigs are particularly distinctive. The fruits of these species are broadly ellipsoid or pear shaped, not compressed, and have dirty whitish or pinkish mesocarp and lanceolate, and brous, non-ligneous leathery endocarp.

Mangifera subsessilifolia shows some af nity with M. lagenifera and M.blommesteinii; however, it has been placed among the species of uncertain taxonomic position due to a lack of complete study material. This is not a very rare species, but owering and fruiting seem to occur at intervals of, or > 5 years, similar in this respect to M. lagenifera, which can be found growing


in old orchards in peninsular Malaysia. The owers and fruits of M. sub-sessilifolia are still unknown.

Mangifera foetida, M. odorata, M. caesia and M. kemanga are widely cultivated in the humid lowlands of the Malay Peninsula, Sumatra, Borneo, Java and Bali. They have also been introduced elsewhere in South-east Asia; M. caesia, M. foe-tida and M. odorata are grown in the southern part of the Philippines, M. foetida is grown in Myanmar, and M. odorata is found in Indochina. They have been described in general reviews of tropical fruit (Ochse and Bakhuizen, 1931; Ochse et al., 1961; Molesworth, 1967; Verheij and Coronel, 1991).

Mangifera pajang, an endemic and commonly cultivated species in Bor-neo, is hardly known outside its native island. This deciduous tree has very stout twigs, with leaves more or less aggregate at the apices. The globose fruits, up to 20 cm in diameter, are the largest known fruits in the genus. The rough, potato-brown rind (0.51 cm thick) can be peeled off like that of a banana. Its bright, deep yellow, thick and brous esh is sweet with a dis-tinctive taste (Kostermans, 1965; Bompard, 1991a). In orchards in Borneo where M. foetida and M. pajang are both cultivated, forms with leaves and fruits having intermediate characters are occasionally found.

Mangifera caesia, M. foetida, M. pajang and especially M. odorata are impor-tant in tropical humid regions where the common mango cannot be grown satisfactorily. Mangifera pajang has potential as an ornamental tree, having brilliant rose-red blossoms (Philipps et al., 1982).

SUBGENUS MANGIFERA. The subgenus Mangifera contains most of the species (47), and is divided into four sections: (i) section Marchandora Pierre; (ii) section Euan-therae Pierre; (iii) section Rawa Kosterm.; and (iv) section Mangifera Ding Hou.

Section Marchandora Pierre. This section has only one species, M. gedebe Miquel (syn. M. camptosperma Pierre, M. inocarpoides Merr. and Perry, M. reba Pierre). The labyrinthine seed is unique to this species, wherein the inner integu-ments penetrate the cotyledons and form numerous irregular folds. The at, discus-like fruit has only a very thin mesocarp. Mangifera gedebe grows in inundated places along rivers or lakes. The seed oats in water and is dis-persed during periods of high water, and this may explain its wide distribu-tion, from Myanmar through Malesia to New Guinea and the Bougainville Island.

Section Euantherae Pierre. The three species in this section (M. caloneura Kurz (syn. M. duperreana Pierre), M. cochinchinensis Engler and M. pentandra Hook. f.) appear to be the most primitive among the species of the subgenus Mangifera. The owers are characterized by the presence of ve fertile sta-mens. The three species are mainly con ned to Myanmar, Thailand, Indo-china and the north of the Malay Peninsula. The region is in the transition zone from the humid tropical rainforest to monsoon forest, and these species show an adaptation to low rainfall. Mangifera cochinchinensis, which occurs in south-eastern Thailand and in Vietnam, has small oblong fruits with a thin seed; the fruits are much relished by local people in southern Vietnam, although they are very acidic. Mangifera caloneura and M. pentandra are closely

J.M. Bompard26

related, and can be mistaken for M. indica. However, their leaves are more leathery, have a more conspicuously dense reticulation, and the panicles are much more hirsute than the common mango. Mangifera caloneura occurs from Myanmar through Thailand to Indochina, in lowland evergreen forests, as well as in semi-deciduous forests. It is cultivated for its acidic-sweet fruit, and has been planted along the streets of Vientiane and Ho Chi Minh City (Saigon). Mangifera pentandra, apparently native to the northern Malay Pen-insula close to the Kra isthmus transition zone, is found in old orchards, in scattered locations, especially in Kedah and possibly also in peninsular Thai-land. It is also grown in the Anambas Islands and in Sabah, where it might have been introduced in early times. It is a proli c bearer, with small man-goes, c.8 cm length, and ripening green or yellow. The pale orange, watery pulp has a sweet taste and few bres.

Section Rawa Kosterm. This group, consisting of nine species, is not well delimitated. Most species have thick twigs and rather coriaceous leaves seated on protruding pedestals. The small, hardly attened ovoid or ellip-soid fruits that are black or partly red at maturity in several species are also characteristic. Rawa is the Malay word for marsh, indicating that these spe-cies usually are found in periodically or permanently inundated areas. The ve species that occur in west Malesia (M. gracilipes, M. grif thii, M. micro-phylla, M. paludosa and M. parvifolia) grow primarily in the swamps of south peninsular Malaysia, in central coastal areas of east Sumatra and western Borneo, and occasionally in peripheral uplands. It has also been reported from the Andaman Islands and from Thailand (Sreekumar et al., 1996; Eiad-thong et al., 2000a).

Mangifera andamanica and M. nicobarica are endemics from the Andaman and Nicobar Islands, respectively. Mangifera merrillii is a rare species endemic to the Philippines and M. minutifolia is known solely from a single collection from southern Vietnam. Mangifera grif thii and M. microphylla are the only cultivated species within section Rawa. The former species is considered to be representative of the section, and is cultivated along the eastern coast of peninsular Malaysia and in western Borneo, and rarely in Sumatra. The fruits are small (35 cm long) and oblong or ovoid; the skin is rose-red, turning purplish black at maturity. The rind is thin and easily removed from the orange-yellow pulp, which is juicy and pleasantly sweet. Different forms are recognized by local people, according to the size and taste of fruits. Mangiferamicrophylla is a related, but less well-known species, having thinner leaves and a rather similar fruit.

Section Mangifera Ding Hou. With more than 30 species, section Mangifera is by far the largest. The common mango and the related M. laurina belong here. Species within the section have the same distribution range as the genus. The section may be divided into three groups based on oral structure and organ number variation: (i) those having pentamerous owers; (ii) those having tetramerous owers; and (iii) an intermediate group of species hav-ing both pentamerous and tetramerous owers. Within these three groups, it is possible to distinguish species with either puberulous or glabrous panicles.


Only characteristics of representative species within each group, especially those found in cultivation, are described below.

Pentamerous owers (14 species): Three species, M. laurina, M. minor and M. sylvatica, show af nity with the common mango. Mangifera laurina is a species of the lowland forests of Malesia, where it is also under cultivation in old orchards. It can be distinguished from the common mango by having lax and widely pyramidal, glabrous or sparingly puberulous panicles. The ow-ers are smaller and are not glomerulate; the petals have a different shape, texture and colour. The fruit resemble those of a small common mango, with orange-yellow pulp, which is almost liquid at maturity. It is generally con-sumed when unripe. Several forms are in cultivation; however these are now becoming rare. Mangifera laurina is well suited to the humid tropical lowlands, fruiting well in areas where the common mango cannot be grown satisfacto-rily; moreover, it appears to be highly resistant to anthracnose (Bompard, 1991b).

Mangifera minor occurs east of Wallaces line, from Sulawesi to New Guinea (east Malesia) and to the Carolines Islands in the east. It is adapted to a wide range of ecological conditions, growing equally well in dry savannahs and in tropical rainforests up to 1300 m. The fruit is obliquely oblong, 510 cm long, much narrowed, the tip obtuse, with a distinct beak and sinus. It is found in cultivation, although the yellowish fruit pulp is acidic and scant. Mangifera sylvatica is found from Sikkim (up to 1200 m) to northern Myan-mar and Thailand, and apparently also in Yunnan up to 1900 m. The fruit is obliquely ovate, 810 cm long, much compressed distally forming a hook, has scanty whitish-yellow pulp which is almost breless. Other species are occasionally found in cultivation, for example M. rufocostata, which is esteemed by the Banjarese people of South Kalimantan for its very sour fruits that are used to prepare a spicy condiment with chilli.

Tetramerous owers (15 species): Mangifera altissima is apparently endemic to the Philippines, where it occurs mainly at low elevations in the forests from northern Luzon to Mindoro (Brown, 1950). The fruit is mango shaped, ovoid or ellipsoid, slightly compressed, up to 8 cm length, green or some-what yellow when ripe, with whitish, sweetish-acidic esh. It is commonly found in dooryards, and thrives in regions with distinct wet and dry seasons (Angeles, 1991).

Mangifera torquenda occurs wild in west Malesia, and is cultivated in south Sumatra and in Borneo, where it is common in the forests and orchards of eastern Kalimantan. The sub-globose fruit, c.7.5 cm long and 6.5 cm in diameter, is yellow-green with darker spots at maturity, and has a thin rind. The pale yellow pulp has a rather pleasant sweet-acid, slightly resinous taste and a light turpentine smell. Short bres are attached to the seed. It is closely related to M. longipetiolata.

Mangifera magni ca is a common species in the rainforests of western Malesia, occasionally cultivated in central Sumatra and in West Kalimantan, where it has a special importance in the myths of Land Dayak peoples. The fruit is ovoid oblong, up to 12 cm long, 10 cm in diameter, only slightly compressed, greyish green with brown spots. The pulp is whitish, soft at

J.M. Bompard28

maturity, sweetish acid. Sweeter forms are reported in central Kalimantan (J.J. Afriastini, personal communication). The stone is unique in the genus in that it lacks bres adhering to it.

Mangifera quadri da is found from peninsular Malaysia to the Moluccas. The fruit is ellipsoid-globose, 68 cm long, green covered with black dots turn-ing completely black at maturity, and has a pale yellow, sweet-acid pulp. Another form is recognized by its more coriaceous leaves, smaller fruits, c.4 cm long, having dark yellow pulp, purplish around the stone, and a sweet, palat-able taste, somewhat like prunes. Both forms are cultivated in old orchards.

Tetra- and pentamerous owers (four species, and also M. indica): Mangifera casturi is related to M. quadri da, from which it can be distinguished by leaf and fruit characters. It has never been collected in the wild, and is a favourite among the Banjarese people in south Kalimantan. The fruits are small, a little compressed and up to 6 cm in length, becoming completely black at matu-rity. The orange pulp is very sweet and palatable, and resembles honey mango or mangga madu grown in East Java. Although M. casturi bears heavily, it has a strong- to alternate-bearing habit. It is an excellent fruit for the humid tropical lowlands, and appears to be resistant to anthracnose. Sev-eral differently named forms exist; these have polyembryonic seeds. Mangiferarubropetala is also only known in cultivation, and may be a primitive race of M. indica.

SPECIES OF UNCERTAIN TAXONOMIC POSITION. There is a group of 11 disparate spe-cies of uncertain taxonomic position that cannot be placed with certainty due to the absence of adequate material. There are three species only known in China.

2.4 Phytogeography

Species distribution

An examination of the present distribution of the genus shows that the larg-est number of Mangifera species in either subgenera is found in western Malesia on the Sunda shelf. A decreasing number of species occurs towards the genus boundary east of Wallaces line in east Malesia, and in its northern and western range of distribution. While peninsular Malaysia and the islands of Sumatra and Borneo have the highest diversity of species, the number of species becomes gradually lower in east Malesia, especially in the Lesser Sunda Islands, Moluccas and New Guinea. This is explained by the geologic and paleogeographic features of the Malesian region which spans two large partly submerged continental shelves, the Asiatic shelf (Sunda Shelf linking the Malay Peninsula with the islands of Sumatra, Java, Borneo and Palawan) and the Australasian shelf (Sahul Shelf linking the Aru islands and New Guinea with Australia). During the last glaciation period (c.22,50011,000 bp) the shelves were regions of land uncovered by the lowering of sea level, and present day peninsular Malaysia, Sumatra and Borneo were connected by


land bridges during the late period of maximal sea lowering. During the cool periods of glacial maxima, the Malesian forest was reduced in extent, but there is no evidence that it was reduced to isolated island forests. The Sunda-land and Papuasian rainforest blocks are therefore comparable to refugia in terms of species richness and the high degree of endemism (Whitmore, 1981). Mangifera has undergone major species development in west Malesia, which has remained relatively stable over a long period of time. The current vegeta-tion of west Malesia probably differs very little from that at the end of the Tertiary (van Steenis, 1950). A lower number of Mangifera species is found in Java and the Philippines, regions less often connected with Asia during the Pleistocene.

Only three species occur in New Guinea, which is largely covered with rainforest. These include M. minor, M. mucronulata and the widely distributed M. gedebe. Mangifera foetida also occurs, but may have been introduced. Mangifera minor occurs from Celebes and the Philippines to the Solomon Islands; M. mucronulata is found in the Moluccas, New Guinea and the Solomon Islands. The distribution of these species suggests a late immigration of a Laurasian genus from Sundaland via the Philippines, Sulawesi and the Moluccas into New Guinea, which is supported by the geological history of the region. No Mangifera species have ever been recorded from northern Australia.

Very few species are found in peninsular India and Sikkim. From present-day distribution, there is little evidence of migration of species into the sub-continent of India after its collision with Eurasia in the middle Eocene (Audley-Charles et al., 1981). According to Mehrotra et al. (1998), fossil leaves described as Eomangiferophyllum damalgiriensis Mehr. from the Upper Palaeo-cene in north-eastern India are an analogue of the modern genus Mangifera.

Mangifera sylvatica occurs along the northern limit of the range of Mangifera, with more or less discontinuity, from Sikkim to northern Thailand and to the southern part of Yunnan, where it is reported in mountains up to 1900 m above sea level (Anonymous, 1980). The few species that grow in southern China are very poorly known: M. austro-yunnanensis from western Yunnan, M. persiciformis from south-eastern Yunnan and southern Guizhou at latitudes up to 26N and M. hiemalis, the winter mango from Guangxi near the northern border Vietnam. In the revised Flora of China (Min and Bar-fod, 2008), M. austro-yunnanensis is considered to be conspeci c with M.indica, M. hiemalis is treated as a synonym of M. persiciformis, and M. laurina is recorded from the lowland forests of south Yunnan.

Subgenera and section distribution

The species distribution is especially meaningful when the ranges of the spe-cies of each subgenus and section are considered separately.

Subgenus LimusAll species of the subgenus Limus are restricted to the Malesian area (M. foetida and M. macrocarpa occurring in peninsular Thailand), whereas all the species

J.M. Bompard30

with ve fertile stamens, considered the most primitive condition, are con- ned to west Malesia (M. decandra in Sumatra and Borneo; M. lagenifera in the two latter areas and in peninsular Malaysia). Only M. caesia, M. foetida and closely related M. leschenaultii occur in east Malesia.

Subgenus MangiferaIn the subgenus Mangifera, M. gedebe is the only species belonging to the sec-tion Marchandora, and has the widest range within the genus, extending from Myanmar through Malesia to New Guinea and Bougainville Island. Section Euantherae is centred in the region from Myanmar to Vietnam. Mangiferapentandra is only known from peninsular Malaysia, the Anambas Islands and Borneo. Section Rawa is mainly in western Malaysia and shows notable diversi cation in the swamps and peripheral uplands in the south of penin-sular Malaysia, east central Sumatra (notably the Riau province) and west Borneo. During the glacial period this area, termed the Riouw pocket (Cor-ner, 1978), formed a vast plain connecting the Malay Peninsula, Sumatra and Borneo, and is believed to have been lled with swamps. Mangifera merrillii is an endemic of the Philippines, M. minutifolia is an endemic of Vietnam, M.andamanica and M. nicobarica are endemics of the Andamans and Nicobar Islands. None of the species of section Mangifera occurring in mainland South-east Asia, north of the isthmus of Kra, are found in eastern Malesia; however, it would be interesting to assess the genetic relatedness of M. syl-vatica and M. minor, and also M. laurina, which may prove to be phylogeneti-cally very closely related.

2.5 Interspeci c Molecular Characterization

Molecular biology techniques now make it possible to assess genetic related-ness in a more precise way. Published data support some of the groupings based on anatomical characters (Kostermans and Bompard, 1993) but not entirely.

RAPD (random ampli cation of polymorphic DNA) markers were rst used in mango by Schnell and Knight (1993) and Schnell et al. (1995). Nine Mangifera species were analysed and compared to the traditional taxonomic groupings. The unweighted pair group method of arithmetic averages (UPGMA) cluster analysis for the subgenus Limus was not supportive of the separation between sections Perennes and Deciduae, which, admittedly, has a weak taxonomic basis. It con rmed the relatedness between M. foetida and M. pajang. The UPGMA cluster analysis of the subgenus Mangifera supported the current taxonomy based on ower morphology. It showed the related-ness between M. quadri da and M. torquenda (both placed in the group of species with tetramerous owers), but also with M. casturi, although the lat-ter species has tetra- and pentamerous owers. One of the most signi cant results was the evidence for the existence of interspeci c hybridization within the studied species of the section Mangifera (see also Yonemori et al., 2002).

Phylogenetic relationships among 14 Mangifera species of Thailand were analysed by comparing ampli ed fragment length polymorphism (AFLP)


markers (Eiadthong et al., 2000b), and by comparing sequences of the inter-nal transcribed spacer (ITS) region of nuclear ribosomal DNA (nrDNA) (Yonemori et al., 2002). They demonstrated that the common mango was closely related to M. laurina, M. sylvatica and M. oblongifolia of subgenus Mangifera to which M. indica belongs. A close relationship between M. indica and M. sylvatica has been corroborated by Nishiyama et al. (2006), who com-pared signal intensity of genomic in situ hybridization (GISH) on somatic metaphase chromosomes of M. indica, using labelled DNA of eight wild Mangifera species.

Furthermore, Eiadthong et al. (2000b) and Yonemori et al. (2002) have demonstrated that M. odorata, M. foetida and M. macrocarpa (of subgenus Limus) were related to M. indica. It is not surprising in the case of M. odorata whose hybrid origin (M. foetida M. indica) has now been established, but this calls into question the position of the section Perennes.

Results of molecular studies do not permit a comprehensive view of the phylogenetic relationships among the genera. So far, they are rather support-ive of the groupings based on phenotype within the subgenus Mangifera (notably sections Rawa and Euantherae), but not for the subgenus Limus which will need to be redescribed, and likely restricted to the group of species related to M. caesia (M. kemanga, M. lagenifera, M. superba and possibly M.decandra). More studies will be needed to infer phylogenetic relationships within the section Mangifera. Keeping in mind the frequent misidenti cations in collections and botanic gardens, herbarium specimens of studied material must be deposited in the national herbaria so that its taxonomic position can be ascertained in case of doubt.

2.6 Region of Origin of the Genus

Based on morphological, phytogeographical and fossil evidence, Mukherjee (1953) argued that:

although the highest number of species of both sections is concentrated in the Malay Peninsula [19 were then recorded], the centre of origin of the genus cannot be restricted to that area alone, as both the phylogenetically older species, i.e. with pentacyclic owers (M. duperreana, now reduced to M.caloneura, and M. lagenifera), occur in Siam and Indochina, and the former is absent from Malay Peninsula.

He concluded that the genus had its origin somewhere in the MyanmarThailandIndochina area or in the Malayan area. Careful identi cation of the greatest part of herbarium materials available has allowed a more accurate delimitation of the distribution ranges of the Mangifera species, notably of the subgenus Limus, and has revealed, among other things, that M. lagenifera does not occur north of Kra isthmus contrary to Mukherjees assertions. Fur-thermore, the ten-stamen species, M. decandra, which was described by Ding Hou in 1972 and hence was unknown to Mukherjee, is con ned to Borneo and Sumatra, and to date has not been recorded from peninsular Malaysia.

J.M. Bompard32

Without overemphasizing the present great species diversity of subgen-era in the Malay Peninsula, Borneo and Sumatra, the available evidence points to a Sundaic origin for the genus. This, however, must not minimize the particular importance of the region stretching from Myanmar to Indo-china as another centre of diversi cation, as attested by the range of species belonging to the section Euantherae. Unfortunately, many of the species of this region remain poorly known, and it can be expected that plant collecting in the region will yield interesting new ndings. The speciation that occurred in this region with a likely radiation centre today traced by the range of the section Euantherae, is of special signi cance as it has given rise to the com-mon mango.

2.7 Origin of the Common Mango

The common mango apparently originated in regions on the western border of the secondary centre of diversi cation mentioned above. Truly wild mango trees have been recorded in Bangladesh (Chittagong Hill tract, c.23N), north-eastern India (undoubtedly indigenous in the evergreen tracts of valley of Assam according to Kanjilal et al., 1937), and in Myanmar where it was reported as not unfrequent in the tropical and lower mixed forests all over Burma from Arracan and Pegu down to Tenasserim (Kurz, 1877). It would be desirable to assess its af nity with the species of the section Euantherae, as well as with species of other sections of the subgenus Mangifera that occur in the same area and region. It is also believed to be wild in the sub-Himalayan tract, in deep gorges of the Baraitch and Gonda hills in Oudh, and the outer hills in Kamaon and Garhwal (Brandis, 1874). The common mango has been grown and disseminated for such a long time in India that semi-wild trees can be found in the forests throughout the subcontinent. The fruits of wild trees are said to be small and of poor quality. Watt (1891) mentioned two so-called almost unaltered wild varieties existed under cultivation in Tirhoot, one originating from Kangra, a very variable one, and the other from Sikkim which was evidently the progenitor of the varieties cultivated in Malda.

The common mango in South-east Asia

The Linnean binomial (Mangifera indica) indicates in this instance the place where the common mango was selected and improved, and not necessarily its place of its origin. It has been traditionally accepted that mango was domesti-cated several millennia ago in India (see Mukherjee and Litz, Chapter 1, this volume); however, it cannot be excluded that domestication occurred inde-pendently in several areas, possibly in the south-western and south-eastern regions of its centre of origin, or later differentiated in those two regions. This hypothesis would account for the differences that exist between the local polyembryonic cultivars of Myanmar, Thailand, Indochina and Indonesia, and the monoembryonic Indian cultivars. Note that polyembryony occurs


also in the cultivated M. casturi, M. laurina and M. odorata. Aron et al. (1998) have demonstrated that polyembryony in mango is under the control of a single dominant gene.

According to Juliano (1937), Bijhouwer suggested that there were two main centres of domestication of mango, one in India with monoembryonic mangoes, the other in the Saigon area, Indonesia and the Philippines with polyembryonic mangoes. The Saigon area must in fact be extended to southern Vietnam, other parts of Indochina, Thailand and Myanmar, which were recognized by Valmayor (1962) as homes of polyembryonic mangoes. Notwithstanding, the origin of polyembryonic mangoes is probably better placed in Myanmar, and possibly the eastern part of Assam. According to Brandis (1874), in Burma, the mango is not generally grafted, and seeds of a good kind, as a rule, produce fruit of a similar description. There are only a few polyembryonic mango cultivars in India. They are restricted to the south-western coastal region, and ge

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